Abiotic Factors (non-soil) and M. peltata Invasion Levels
BIOTROPIA Vol. 23 No. 1, 2016: 21 - 27 DOI: 10.11598/btb.2016.2 . . 3 1 457
ABIOTIC FACTORS INFLUE NCING MANTANGAN
( Merremia peltata ) INVASION IN BUKIT BARISAN SE LATAN
1*NATIONAL PARK
2 3 JANI MASTE R , SOE KISMAN TJITROSOE DIRDJO and IBNUL QAYIM 1 Department of Biology, Faculty of Mathematics and Natural Sciences,Universitas L ampung, Bandar L ampung 35145, Indonesia
2 3 Weed Science Society of IndonesiaDepartment of Biology, Faculty of Mathematics and Natural Sciences,
Institut Pertanian Bogor, Bogor 16680, Indonesia
Received 20 January 2015/Accepted 27 January 2016
ABSTRACT
Some areas in Bukit Barisan Selatan National Park have serious problems related to Merremia peltata (Family Convolvulaceae) invasion. The abiotic factors influencing M. peltata invasion were investigated in this study. This research compared abiotic factors in three forest locations with different invasion levels i.e. severe, moderate and mild invasions. Abiotic factors measured were percentage of tree canopy coverage, micro climate factors (temperature and humidity), elevation above sea level and physical and chemical properties of the soil surface. In this study, canopy coverage is classified as abiotic factors because it influences the intensity of light that can reach the forest floor. A post- hoc Duncan's Multiple Range Test (DMRT) was employed to determine significant differences ( < 0.05) among p abiotic factors. In addition, a test of correlation and multiple linear regression were conducted to determine the relationships between abiotic factors and M. peltata invasion. Multiple regression testing showed that canopy coverage significantly ( <0.05) influenced p M. peltata invasion. Based on the generated model, a 1% addition of canopy coverage would decrease M. peltata invasion by 2.8% Multiple linear regression analysis was used to predict the relationship . between soil nutrients and invasion. The C/N ratio and P O , Ca, Mg and Na concentration significantly ( < 0.05) p 2 5 influenced invasion level. Again, based on the generated model, the increase in the C/N ratio as well as in P O and Ca 2 5 concentrations were correlated with the reduction of M. peltata invasion; while the increase of Mg and Na concentrations were correlated with the increase of M. peltata invasion. M. peltata invasion in Bukit Barisan Selatan National Park was influenced by opened forest canopy and was correlated with high concentration of Na and Mg in the soil of the invaded forest areas.
Keywords : Bukit Barisan Selatan National Park, invasive species, mantangan, Merremia peltata
INTRODUCTION economy, environment and/or cause damage to
human, animal or plant health (National Invasive Indonesia is a country with abundant flora and Species Council 2006). fauna, some of which are conserved and Merremia peltata or mantangan in the local protected in conservation areas such as national language, is an invasive species of conservation parks. E xisting national park areas in Indonesia areas. This species causes severe problems in are threatened by alien plant species that invade Bukit Barisan Selatan National Park (BBSNP) and threaten flora and fauna biodiversity. with more than 7,000 ha covered by M. peltata
Alien species that have the ability to grow (Master 2013). Infestation of the weed is et al. quickly so as to impact biodiversity in invaded suspected to be the cause of animal migrations, areas can be referred to as Invasive Alien Species such as tiger, elephant and Sumateran rhinoceros, (IAS) (Pyšek & Richardson 2010). Broadly, an IAS to rural areas up north (Irianto & Tjitrosoedirdjo is a species that has the potential to harm the 2010). Analysis of Landsat and Advanced Land
Observing Satellite (ALOS) images indicated that
- C orresponding author: j.janter@ gmail.com
M. peltata invasion is found from south part of BBSNP running 33 km to the north of the national park. It is also found in small outbreaks as far as 50 km north of BBSNP (Master 2013). et al.
The BBSNP is the third biggest conservation area in Sumatera Island covering an area of 356,800 ha. Administratively, this area belongs to Lampung Barat, Pesisir Barat and Tanggamus Districts of Lampung Province and the Bengkulu District of Bengkulu Province, (4 31' – 5 57' S; 103 34' – 104 43' E ) (Gaveaua 2007). et al.
M. peltata is a liana (climbing plant) of the
Convolvulaceae et al. family (Paynter 2006), and it is distributed throughout Madagascar, the Mescarenes, Seychelles, Malay Peninsula, Malaya Islands, Philippines, New Guinea, Northen Aus t r al i a an d t h r o ugh o ut P o l yn e s i a (Ooststroom & Hoogland 1953). It is similar to sweet potato in morphology, with a wide heart shaped leaf connected by peltate petiole, in contrast to the petiole connection of sweet potato. The hairless plant stem grows to 20 m in length and excretes white liquid when injured. It may grow over other plants (Stone 1970). The corolla is white or yellow, 5–6 cm in length and forms a bell-shape corolla (Fosberg & Sachet 1977). Spreading of this plant occurs generatively, both by seeds or vegetatively from roots along its stems which touch the ground. egetative growth
V rates reach 16 cm/week can (Pengembara 2014).
E nvironmental factors, both biotic and abiotic, determine the presence of a species in a particular location (Huang 2003). et al. Accordingly, not all alien species introduced to an area will become invasive; this can be influenced by factors including environmental fitness and the presence or absence of predators or competitors (Theoharides & Dukes 2007). The rapid growth of in BBSNP is probably caused by a M. peltata number of abiotic factors. The objective of this research was to determine abiotic factors influencing invasion in BBSNP M. peltata .
M erremia peltata is found mostly in southern
regions of BBSNP, but not all of in this M. peltata region grows densely. To determine abiotic factors influencing plant invasion, environmental factors were recorded in three forest locations with different invasion levels, i.e. severely invaded, moderately invaded and mildly invaded forest (Fig. 1). First location (severely invaded forest) is close to the village enclave Way Haru, Bandar Dalam and Tampang. Based on land cover maps, this site has a decreasing forest cover for an area of 2,565.54 ha from 2000 to 2009. Presumably these changes were the result of forest clearing
BIOTROPIA Vol. 23 No. 1, 2016 Figure 1 Map of research location of invasion in Bukit Barisan Selatan National Park, Lampung, Sumatera M. peltata (Master 2013) et al.
MATE RIALS AND ME TH ODS
Abiotic factors influencing mantangan ( ) invasion Jani Master Merremia peltata – et al. for plantations (Prasetyo 2011). The second et al. location (moderately invaded forest) is a post-fire forest in 1997; fires that occurred mostly on the forest floor affecting the lining of plant and open up some forest canopy (WCS-IP 2001). The third location (mildly invaded forest) is a primary forest. Selection and classification of these invasion levels were based on coverage of from
Data for each environmental factor from each of the three locations were analyzed using ANOVA and the mean differences were tested using the Duncan's Multiple Range Test (DMRT) at < 0.05). Tests of correlation and linear p regression were carried out using SPSS 12 program to determine any relationships existed between abiotic factors and invasion.
Landsat and ALOS image analyses in 2002 and 2008 (Master 2013). The final selection was et al. conducted by recording the percentage of
M. peltata
M. peltata
RE SULTS AND DISCUSSION
(CE C) and exchangeable cations (K, Na, Ca, Mg). Soil texture fraction comparisons were separated as follows: 1. sand fraction having particles with diameter of 2 mm - 50µ; 2. loam fraction having particles with diameter of 50µ - 2µ; and 3. clay fraction having particles with diameter of < 2µ) (Hardjowigeno 2010).
Ground temperature ( o
Note: Numbers followed by the same letters in same columns are not significantly different ( < 0.05) p
54.05 a 6.51 a 69.89 c
27.15 ab 88.84 a 26.00 ab
27.93 b 83.79 b 26.10 b 47.65 a 6.65 a 81.51 b Severely invaded forest
Mildly invaded forest 26.41 a 90.72 a 25.65 a 47.93 a 6.75 a 92.31 a Moderately invaded forest
Canopy coverage (%)
(%) Soil pH
C) Soil humidity
C) Air humidity (%)
canopy coverage by using plot of 20 x 20 m area. Three line transects of 1,000 m each, with minimum of 200 m interval distance between each line transect, were made in each forest location. Plots of 20 x 20 m were sampled along each line transect, at 100 m intervals, thus totaling 30 plots in each location. invasion was M. peltata estimated based on the percentage of plot that was covered by . Environmental factors M. peltata recorded for each plot were: 1. the percentage of tree canopy coverage (tree canopies that were more than 4 m in height) which was estimated using a concave densiometer; 2. microclimate (temperature and humidity) were measured at noon, 1 m above the ground using thermometer and hygrometer on each plot; 3. elevation above sea level; and 4. physical and chemical properties of top soil using composite method (Crozier . et al 2010). Soil physical and chemical properties observed were soil texture, pH (H O), KCl, C- 2 organic, N-Kjedahl, Cation E xchange Capacity
Air temperature ( o
Table 1 Summary of abiotic factors at the three research locations Location
M. peltata
peltata invasion by 2.8% (Table 2).
Of the six abiotic factors observed i.e. air temperature, air humidity, ground temperature, soil humidity, soil pH and percentage of tree canopy coverage (canopy above 4 m), only the percentage of tree canopy coverage was significantly varied ( < 0.05) among the three p locations (Table 1). Forest canopy coverage had significant negative correlation to M. peltata invasion. Based on a generated model, every 1% addition of canopy coverage would decrease M.
Abiotic Factors (non-soil) and M. peltata Invasion Levels
The mean coverage percentage of in the mildly invaded forest ranged from 1 to 15%. In comparison, the mean coverage percentage in the moderate invaded forest ranged from 27 to 55%. In the severe invaded forest, the mean coverage percentage was 44%, while in some location the percentage was up to 100%.
Percentage Cover of Invading M. peltata Forests M. peltata
The severely invaded forest locations had more open tree canopy coverage. The open canopies facilitate access of to solar M. peltata radiation such that the weed dominated these BIOTROPIA Vol. 23 No. 1, 2016 Table 2 Results of multiple linear regression between percentages of M. peltata coverage and abiotic factors a
Coefficients Standardized
Unstandardized coefficients coefficients Collinearity statistics t
Model Sig.
Std. B
Tolerance
VIF Beta
E rror 1 (Constant) 89.020 53.253 1.672 0.099
Air temperature -2.841 3.189 - 0.148 -0.891 0.376 0.356 2.812 0.100 0.345 2.897
Air humidity - 0.281 -1.668
- 2.782 1.668
0.434 2.317 Ground temperature 6.209 0.119 0.787 0.432
7.892
- 0.236 0.123 Soil humidity
- 1.562 0.431 2
- 0.603 0.386
2.418 Soil pH -23.790 - 0.448 -2.910 0.005 0.414
8.175
- 0.427 0.000 Canopy coverage
0.882
- 1.218 0.301 -4.047
1.134
Note: a = Dependent Variable: percentage of M. peltata coverage
areas. In the mildly invaded forest, mean canopy different to mildly invaded forest. This is coverage was denser (92.31%). Denser canopy probably because the ground coverage of coverage prevented extensive M. peltata growth M. peltata in both levels of invaded forest and development such that it only grew through produced similar microclimate. gaps of canopy where there were collapsed trees. There were no relationship between elevation This result is consistent with other findings which above sea level and the infestation size of reported that M. peltata grew on the edges of M. peltata. This is despite the fact that M. peltata is forest, especially in degraded land in Fiji (Kirkham known to occupy lowland habitats to 300-400 m 2005), and that this plant grew densely in logged above sea level in Samoa and Fiji (Meyer 2000, forest and open forest in other parts of Sumatera Kirkham 2005). (Irianto & Tjitrosoedirdjo 2010).
Air temperature, air humidity and ground Soil Abiotic Factors and M. peltata Invasion temperatures were all significantly ( < 0.05) higher p
Levels
in moderately invaded forest than in mildly Beside the aforementioned environmental invaded forest (Table 1). The increase in these factors, the physical and chemical properties of factors was most likely related to tree canopy the soil were also analyzed in the three research coverage, such that more open canopies would locations. T he mildly invaded forest was increase solar radiation, air and ground dominated with a sandy-clay-loam soil texture, temperatures, as well as rainfall reaching the forest while the moderately and severely invaded forest floor (Sasaki & Mori 1981). had clay and clay-loam soil texture (Table 3). Severely invaded forest had more open tree
Soil in the secondary and invaded forests had canopy coverage, but air temperature, air humidity higher clay content than soil in the primary forest. and ground temperatures in that location were not
The clay fraction is the most important Table 3 Soil texture samples in the three research locations
Location Texture Percentage (%)
Mildly invaded forest sandy clay loam
33.33 clay 55.56 clay loam
11.11 Moderately invaded forest 44.44 clay 55.56 clay loam
55.56 Severely invaded forest clay clay loam
44.44 component because its particles have more surface area than sand particles. Soil with higher clay content has higher Cation E xchange Capacity (CE C) than sandy soil due to greater overall surface area of clay particles (Munawar 2011). The CE C value was higher in the severely invaded forest (25.67 cmol /kg), followed by moderately c invaded forest (19.07 cmol /kg), with the lowest in c the mildly invaded forest (13.51 cmol /kg) (Table c
4). The CE C is a chemical property closely related to soil fertility. Soil with higher organic and clay contents had higher CE C values than soil with low o r ganic c o nt e nt an d/o r sandy so il s (Hardjowigeno 2010).
Very Low Ca (cmol c /kg) 4.59 a
Notes: CE C = Cation Exchange Capacity Numbers followed by the same letters in the same row are not significantly different ( < 0.05) p
CE C * (cmol c /kg) 13.51 a Low 19.07 a Middle 25.67 b High Base saturation (%) 62.55 a High 55.11 ab High 71.11 b High
Middle 0.46 b Middle 0.51 c Middle
0.67 a High Na (cmol c /kg) 0.41 a
K (cmol c /kg) 0.57 a Middle 0.50 a Middle
High 3.96 a High 6.28 b High
Middle Mg (cmol c /kg) 3.04 a
Low 5.50 a Low 11.11 b
Very Low 6.53 b
Further, soil with higher CE C values is able to better retain and provides nutrients (cations and anions) than soil with low CEC values. The lack of cation exchange capacity in low CE C soil results in the nutrients being easily washed through the soil with underground water movement (infiltration, percolation) such that these nutrients are no longer available for plant growth.
K 2 O (mg/100 g) 28.33 a Middle 31.11 a Middle 39.77 a Middle P-Bray (ppm) 4.00 a Very Low 2.76 a
C-organic (%) 2.07 a Middle 2.62 ab Middle 3.46 b High N (%) 0.17 a Low 0.21 a Middle 0.35 b Middle C/N Ratio 11.66 a Middle 11.88 a Middle 9.88 b Low P 2 O 5 (mg/100 g) 24.22 a Middle 18.44 b Low 24.88 a Middle
Mildly invaded forest Moderately invaded forest Severely invaded forest Value Level Value Level Value Level
Table 4 Average soil chemical properties in three research locations Soil chemical property
Double linear regression analyses were conducted to predict whether soil nutrient status was correlated with invasion. C/N ratio and concentration of P O , Ca, Mg and Na 2 5 significantly ( < 0.05) contributed to p M. peltata invasion (Table 5). Based on a generated model, the increase of C/N ratio and concentration of P O and Ca may reduce invasion; while 2 5 M. peltata the increase of Mg and Na concentration may increase invasion. invasion in M. peltata M. peltata the severely invaded forest was correlated with
Regression Analyses between Soil Factors and Invasion M. peltata
Nitrogen (N) is a crucial soil nutrient, often found with low availability. Total nitrogen in the severely invaded forest was significantly ( < 0.05) p higher (0.35%) than that in the other two locations. There was no significant difference ( > 0.05) of total nitrogen between the mildly p invaded forest (0.17%) and the moderately invaded forest (0.21%). Higher nitrogen values contribute to low C/N ratios such that more nitrogen is available to microorganisms to decompose carbon containing material, recycling the nitrogen quickly (Munawar 2011). Higher nitrogen contents in plants will result in faster vegetative growth. was rarely found to M. peltata flower in severely invaded forest, and this may have been caused by high nitrogen contents in the soil.
The presence of organic substances in the soil can improve soil fertility chemically, physically and biologically. The amount of organic carbon in the severely invaded forest was higher (3.46%) than that in the mildly invaded forest. Based on soil fertility criteria, C-organic in the severely invaded forest belonged to high category.
The severely invaded forest had a higher alkaline density (base saturation) value (71.11%) than the other two forest locations. Contributing to this higher alkaline density were the alkaline cations Ca, Mg, K and Na. These alkaline cations are important soil nutrients required by plants.
Abiotic factors influencing mantangan ( ) invasion Jani Master Merremia peltata – et al.
BIOTROPIA Vol. 23 No. 1, 2016 Table 5 Result of multiple linear regression between the percentage of M. peltata coverage and soil nutrient factors a
Coefficients Standardized
Unstandardized coefficients Collinearity statistics coefficients Model t Sig. Std. B Beta Tolerance
VIF E rror 1 (Constant)
50.768 144.015 0.353 0.729 0.315
Loam 0.797 1.181 0.675 0.509 3.173 0.117
0.190 Clay -0.569 0.905 - 0.140 -0.629 0.538 5.277
- 0.128 H 2 O -11.183 23.641 -0.473 0.642 0.128 7.818 CN Ratio -8.535 -3.116 1.498
2.739 - 0.370 0.006 0.668 P O -2.721 1.197 -2.273 2.924 2 5 - 0.377 0.036 0.342
Ca -5.902 1.694 - 0.888 -3.485 0.145 6.899 0.003
Mg 18.726 4.721 0.970 3.966 0.001 0.157 6.351 Na 210.264 74.615 0.390 2.818
0.012 0.493 2.029
Base 0.128 0.284 3.522
0.735 0.292 1.601 saturation (%) 0.459
Note: a = Dependent Variable: coverage of Merremia peltata
low C/N ratios and high concentration of Mg and CONCLUSIONS Na in the soil.
Sodium has an important role in determining Abiotic factors influencing M. peltata invasion soil characteristics and plant growth, especially in in Bukit Barisan Selatan National Park are opened areas close to a beach. The severely invaded forest forest canopy coverage and high content of Na in this study is located close to a beach. This is a and Mg in the soil. suspected cause of the high Na content in the soil in this location. E xcessive Na content can also be toxic for plants (Hanafiah 2007), influencing cell ACKNOWLE DGE ME NTS membrane and organelle functions, as well as disrupting plant metabolism sequences (Hamim This study was part of Master thesis of the 2008). Soil sodium concentrations significantly first author, financially supported by SE AME O- ( < 0.05) increased as invasion levels of p M. peltata BIOTROP DIPA 2011. The author thanked increased in this study. This weed is suspected of Artha Graha Peduli and Wildlife Conservation being a halophyte, being able to grow in a land Society Indonesia Program for their assistance in with high salt content. conducting the field work.
Another soil nutrient correlated with M. peltata invasion was Mg. Soil samples from the severely
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